Permafrost microbial communities and functional genes are structured by latitudinal and soil geochemical gradients.

Permafrost underlies approximately one quarter of Northern Hemisphere terrestrial surfaces and contains 25-50% of the global soil carbon (C) pool. Permafrost soils and the C stocks within are vulnerable to ongoing and future projected climate warming. The biogeography of microbial communities inhabiting permafrost has not been examined beyond a small number of sites focused on local-scale variation. Permafrost is different from other soils. Perennially frozen conditions in permafrost dictate that microbial communities do not turn over quickly, thus possibly providing strong linkages to past environments. Thus, the factors structuring the composition and function of microbial communities may differ from patterns observed in other terrestrial environments. Here, we analyzed 133 permafrost metagenomes from North America, Europe, and Asia. Permafrost biodiversity and taxonomic distribution varied in relation to pH, latitude and soil depth. The distribution of genes differed by latitude, soil depth, age, and pH. Genes that were the most highly variable across all sites were associated with energy metabolism and C-assimilation. Specifically, methanogenesis, fermentation, nitrate reduction, and replenishment of citric acid cycle intermediates. This suggests that adaptations to energy acquisition and substrate availability are among some of the strongest selective pressures shaping permafrost microbial communities. The spatial variation in metabolic potential has primed communities for specific biogeochemical processes as soils thaw due to climate change, which could cause regional- to global- scale variation in C and nitrogen processing and greenhouse gas emissions.

Predicting Effects of Ocean Acidification and Warming on Algae Lacking Carbon Concentrating Mechanisms.

Seaweeds that lack carbon-concentrating mechanisms are potentially inorganic carbon-limited under current air equilibrium conditions. To estimate effects of increased atmospheric carbon dioxide concentration and ocean acidification on photosynthetic rates, we modeled rates of photosynthesis in response to pCO2, temperature, and their interaction under limiting and saturating photon flux densities. We synthesized the available data for photosynthetic responses of red seaweeds lacking carbon-concentrating mechanisms to light and temperature. The model was parameterized with published data and known carbonate system dynamics. The model predicts that direction and magnitude of response to pCO2 and temperature, depend on photon flux density. At sub-saturating light intensities, photosynthetic rates are predicted to be low and respond positively to increasing pCO2, and negatively to increasing temperature. Consequently, pCO2 and temperature are predicted to interact antagonistically to influence photosynthetic rates at low PFD. The model predicts that pCO2 will have a much larger effect than temperature at sub-saturating light intensities. However, photosynthetic rates under low light will not increase proportionately as pCO2 in seawater continues to rise. In the range of light saturation (Ik), both CO2 and temperature have positive effects on photosynthetic rate and correspondingly strong predicted synergistic effects. At saturating light intensities, the response of photosynthetic rates to increasing pCO2 approaches linearity, but the model also predicts increased importance of thermal over pCO2 effects, with effects acting additively. Increasing boundary layer thickness decreased the effect of added pCO2 and, for very thick boundary layers, overwhelmed the effect of temperature on photosynthetic rates. The maximum photosynthetic rates of strictly CO2-using algae are low, so even large percentage increases in rates with climate change will not contribute much to changing primary production in the habitats where they commonly live.

Reproductive effort of Mastocarpus papillatus (Rhodophyta) along the California coast.

Species with sexual and asexual life cycles may exhibit intraspecific differences in reproductive effort. The spatial separation of sexual and asexual lineages, called geographic parthenogenesis, is common in plants, animals and algae. Mastocarpus papillatus is a well-documented case of geographic parthenogenesis in which sexuals dominate southern populations, asexuals dominate northern populations, while mixed populations occur throughout central California. We quantified abundances and reproductive effort of sexual and asexual fronds and tetrasporophytes at eight sites in California to test the hypotheses that (1) reduced sexual reproduction at higher latitudes and tidal heights explains the observed geographic parthenogenesis and (2) reproductive effort (spore production per blade area) declines with increasing latitude. Abundances of all phases varied site-specifically. However, there was no geographic pattern of reproductive effort of fronds. Reproductive effort of fronds was greater in 2006 than in 2007 and correlated with sea surface temperatures. Sexual fronds exhibited greater reproductive effort than did asexual fronds and sexual reproductive effort was also inversely correlated with local upwelling index. Tetrasporophytes showed greater reproductive effort in northern sites, but total supply of tetraspores per m2 was greatest in the middle of the sampling range where crusts were more abundant. There was no decline of reproductive effort at higher latitudes. Geographic patterns of fecundity of life stages do not explain geographic parthenogenesis in M. papillatus. Site-specific differences in viability among spores or established thalli of different life cycles may explain their respective geographic distributions, as the sexual and asexual life cycles responded differently to environmental variations.

Spatial distribution and reproductive phenology of sexual and asexual Mastocarpus papillatus (Rhodophyta).

Species of the genus Mastocarpus exhibit two distinct life cycles, a sexual alternation of generations and an obligate, asexual direct life cycle that produces only female upright fronds. In the intertidal red alga, M. papillatus (Kützing) sexual fronds dominate southern populations and asexual fronds dominate northern populations along the northeast Pacific coast, a pattern of spatial separation called geographic parthenogenesis. Along the central coast of California, sexual and asexual variants occur in mixed populations, but it is not known whether they are spatially separated within the intertidal zone at a given site. We investigated reproductive phenologies and analyzed patterns of spatial distributions of sexual and asexual M. papillatus at three sites in this region. Sexual M. papillatus were aggregated lower on the shore at two sites and only reproduced during part of a year, while asexual M. papillatus occurred throughout the intertidal range at all sites and reproduced throughout the year. The distribution patterns of sexual and asexual M. papillatus are consistent with a hypothesis of shoreline topography influencing their dynamics of dispersal and colonization. Spatial and temporal partitioning may contribute to the long-term coexistence of sexual and asexual life histories in this, and other, species of Mastocarpus. The occurrence of geographic parthenogenesis at multiple spatial scales in M. papillatus provides an opportunity to gain insight into the phenomenon.